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밀 잎녹병 저항성 유전 육종 주요 연구 동향

김경민1, 조성우2, 강천식1, 김경훈1, 최창현1, 손재한1, 박철수3, 모영준1,*

Research Advances in Wheat Breeding and Genetics for Leaf Rust Resistance

Kyeong-Min Kim1, Seong-Woo Cho2, Chon-Sik Kang1, Kyeong-Hoon Kim1, Chang-Hyun Choi1, Jae-Han Son1, Chul Soo Park3, Youngjun Mo1,*

Published online: September 1, 2020

1국립식량과학원

2경남과학기술대학교 농학한약자원학부

3전북대학교 작물생명과학과

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*Corresponding Author (E-mail: moyj82@korea.kr, Tel: +82-63-238-5232, Fax: +82-63-238-5205)
• Received: May 6, 2020   • Revised: May 7, 2020   • Accepted: May 22, 2020

Copyright © 2020 by the Korean Society of Breeding Science

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  • AbstractExtensive research has been conducted in wheat to improve genetic resistance to rust, a major disease that deteriorates wheat yield and quality worldwide. Leaf rust caused by Puccinia triticina is the most prevalent among the three major wheat rust diseases (leaf, stripe, and stem rust) globally and is adapted to a wide range of climates. Approximately 80 genes for leaf rust resistance have been reported, and six (Lr1, Lr10, Lr21, Lr22a, Lr34, and Lr67) have been cloned. Among these cloned genes, Lr34/Yr18/Sr57/Pm38 and Lr67/Yr46/Sr55/Pm46 are of special interest for breeding programs, as they provide pleiotropic resistance to stripe rust, stem rust, and powdery mildew as well as leaf rust. In Korea, knowledge on wheat breeding and genetics for rust resistance is insufficient, as agronomic measures have mainly been used to avoid rust. Therefore, an extensive research program to address the increasing threat of rust epidemics due to climate change is urgently required. Major Korean wheat cultivars, breeding lines, and landraces should be screened for leaf rust resistance. Diverse germplasms also need be introduced through international collaborations to broaden the genetic background for resistance. It is equally important to characterize the distribution of different leaf rust races in Korea and respond to changes in pathogen populations by using effective resistance genes in breeding programs. Research on molecular genetics and genomics needs to be furthered to identify new leaf rust resistance genes and develop efficient molecular markers.
세계적으로 밀 생산 증대와 품질 향상을 위한 연구가 꾸준히 진행되고 있지만, 경작지 감소와 기후변화, 예측 불가능한 비생물적 및 생물적 스트레스의 위협은 증가하고 있다(Figueroa et al. 2018). 생물적 스트레스인 곰팡이, 바이러스 및 박테리아에 의해 유발되는 질병은 밀의 수량과 품질에 치명적인 결과를 초래한다. 그 중 곰팡이 병원균은 바이러스나 박테리아에 비해 환경 적응력과 변형을 통해서 독성을 빠르게 강화시키고 널리 퍼지게 되는데, 대표적인 곰팡이병인 녹병(rust)은 전세계 밀의 안정적 생산에 가장 큰 위협이 되는 요인 중 하나로, 갈색 반점을 띄는 잎녹병(leaf rust), 노란색 줄무늬를 나타내는 줄녹병(stripe rust 또는 yellow rust), 검은 반점을 보이는 줄기녹병(stem rust 또는 black rust)이 있다(Cummins & Hiratsuka 2004, Duplessis et al. 2011). 그 중 잎녹병은 다른 녹병에 비해서 빈번하고 광범위하게 발생되어 발병으로 인한 수량 감소가 최대 50%에 이르며 피해 규모나 액수가 가장 크다.
밀 잎녹병균인 Puccinia triticina Eriks. (Pt)는 Fertile Crescent 지역이 기원지로 온화한 기온과 습한 조건에서 널리 퍼지는 특성이 있어 녹병중에서 전세계적으로 가장 많이 발생하는 병이다(Bolton et al. 2008, Huerta-Espino et al. 2011). 잎녹병은 병원체가 매우 다양하고 새로운 병원성 양상(virulence profiles)이 지속적으로 나타나며 광범위한 기후에 높은 적응력을 지닌다(Kolmer 2005, Huerta-Espino et al. 2011, McCallum et al. 2016). 미국의 경우 2000년부터 2004년까지 잎녹병으로 약 3억5천만달러의 피해를 보았다(Huerta-Espino et al. 2011, Kolmer 2005). 중국에서는 잎녹병으로 인해 연간 3백만톤의 수량 감소가 보고되고 있는데, 1970년대에는 네 차례의 잎녹병 대발생이 있었고, 밀 주요 재배지역인 북부지역과 황허(黃河)와 화이허(淮河)의 남부지역은 여전히 잎녹병 발생 위험이 커지고 있으며, 다른 여러 지역에서도 잎녹병으로 인한 수량 감소가 보고되고 있다(Zhao et al. 2008, Zhou et al. 2013). 멕시코와 서아시아 지역의 경우 과거 잎녹병 발생으로 수량 감소가 심각하여 멕시코는 1976/7년에 발생한 잎녹병으로 70%까지 수량 감소가 있었으나, 최근 발병 진행이 느린(slow rusting) 저항성 품종이 보급되면서 피해가 다소 줄어들고 있다(Dubin & Torres 1981, Huerta-Espino et al. 2011).
녹병 저항성 유전자는 밀과 밀 야생종으로부터 200여종이 보고되었으며 대부분이 종 특이적(race-specific) 저항성 유전자로 알려져 있다(Periyannan et al. 2017, Aktar-Uz-Zaman et al. 2017). 녹병에 대한 유전적 저항성 확보를 위해서는 전세계 밀 품종 및 야생밀을 포함한 유전자원으로부터 효과적 저항성 유전자를 동정하는 것이 필수적인데, 이러한 작업은 시간과 노력을 많이 요하는 힘든 과정이며, 다수의 저항성 유전자가 뒤섞여 있는 조건에서 단일 저항성 유전자를 탐색하고 동정하기는 더욱 어렵다(Gold et al. 1999, Sumikova & Hanzalova 2010, Kadkhodaei et al. 2012). 또한 녹병균은 일반적 감염수준에서 ha당 2.5×1012 개 가량의 포자를 생성하기 때문에 저항성 품종을 개발하여도 돌연변이에 의해 독성을 새로 획득한 병원균이 지속적으로 발생한다(Fetch et al. 2011). 그러므로 저항성 품종 재배면적이 늘어남에 따라 병원균의 독성도 증가하는 것이 일반적이다. 따라서 단일 저항성 유전자로는 녹병을 극복할 수 없기 때문에 저항성 증진을 위하여 다수의 저항성 유전자의 집적(gene pyramiding)이 필요하다(Mundt 2018). 이를 위해 종 특이적(race-specific) 주동 유전자(major gene)와 종 비특이적(race- non-specific)인 미동 유전자(minor gene)를 분자 표지를 활용하여 품종에 4~5개 정도 집적하는 방법이 제안되고 있다(Periyannan et al. 2017).
잎녹병과 관련된 저항성 유전자는 현재까지 80여개가 보고 되었다(McIntosh et al. 1995, 2013, Dakouri et al. 2013, Qureshi et al. 2018). 잎녹병 저항성 평가에 있어서 유전자 간 상호작용과 환경 영향으로 인하여 평가의 어려움이 있지만, 분자표지를 활용한 MAS (marker-assisted selection) 육종을 통하여 효과적인 저항성 유전자를 집적하는 것이 최상의 대안으로 여겨진다(Periyannan et al. 2017, Mundt 2018). 최근 국내에서도 녹병 발생 횟수가 많아지고 있지만 관련 연구나 정보가 미흡하여 국내 밀의 안정 생산을 위한 녹병 저항성 품종 개발이 시급한 실정이다. 본 논문은 국내 환경에 적합한 잎녹병 저항성 밀 품종 육성에 필요한 정보를 정리하였다.
잎녹병 생활사
녹병균은 성장과 번식을 숙주 세포에 의존하는 기생 영양체(biotrophic organism)로 종류에 따라 특정 숙주를 감염시키는 능력도 다르다(Cummins & Hiratsuka 2004, Duplessis et al. 2011). 잎녹병은 Basidiomycete 과 Puccinia 속의 P. triticina (Pt)가 병을 일으킨다(Bolton et al. 2008). 녹병은 무성생식과 유성생식, 숙주의 종류에 따라서 5가지 형태의 포자를 형성하는데, 여름 포자(urediniospores), 겨울 포자(teliospores)와 담자 포자(basidiospores)는 밀에서 발생하고, 병포자(pycniospores)와 녹포자(aeciospores)는 밀 이외 대체 숙주(alternative host)에서 발생한다(McIntosh 2009, Jin et al. 2010, Chen et al. 2014). 잎녹병의 대체 숙주는 Thalictrum, Anchusa, Clematis와 Isopyrum 속으로 알려져 있다(Leonard & Szabo 2005, Huerta-Espino et al. 2011, Wang & Chen 2017). 대체숙주에서 발달된 포자가 저항성이 없는 밀을 감염시키는데, 잎녹병의 2핵체 여름 포자(dikaryotic urediniospores)는 포자가 수천 km를 갈수 있기 때문에 한번 발생으로 대륙 전체가 문제가 될 수 있어 세계적 수준의 방제 프로그램이 반드시 필요하다(Khan et al. 2013). 잎녹병균은 포자의 발아 과정에서 높은 습도가 요구되며 적정 온도는 15~20°C이고 분생자형성(sporulation)이 되기 전까지는 밀에서 증상이 나타나지 않는다(Bolton et al. 2008). 따라서 우리나라는 비가 자주 오고 기온이 낮아지는 시기에 녹병이 주로 발생하는데, 5월 등숙기와 6월 수확기 경에 특히 많이 발생한다.
잎녹병에 감염되면 주로 잎에 원형이나 타원형의 주황색 농포인 여름 포자가 나타난 후에 점점 검정색의 겨울 포자로 발전하며, 영(glume)과 까락에서도 발병한다(Roberson & Luttrell 1987, Da-Silva et al. 2012, Fig. 1). 잎녹병 감염으로 종실 무게와 1수립수 감소로 수량 감소가 일어나는데, 이러한 피해는 발생 시기와 밀의 생육 단계 및 품종의 저항성 정도에 따라 다양하게 나타나며 분얼기 전 생육초기에 발병 할 경우 막대한 수량 감소가 발생하는 반면 출수 후 발병은 수량 감소 영향이 적다(McIntosh et al. 1995, Bolton et al. 2008).
잎녹병 저항성 기작
두번째 형태는 종비특이적(race-non-specific) 저항성으로, 주로 유묘기 이후에 저항성을 나타내기 때문에 성체 저항성(APR, adult plant resistance)이라고도 한다(Periyannan et al. 2017). APR은 녹병균 잠복기간이 길고 감염 빈도가 낮으며 여름 포자의 크기가 작고 포자 생존 시간도 짧아 감염 부위당 포자 생성이 적은데, 이로 인하여 병도 느리게 발현(slow rusting)한다(Schnurbusch et al. 2004, Singh et al. 2005). 이러한 형태의 저항성 유전자는 대표적으로 Lr34, Lr46, Lr67, Lr68 등이 있지만 이들의 정확한 저항성 기작에 대해서는 알려진 것이 별로 없다(Ellis et al. 2014). 동정이 이루어진 Lr34, Lr67에서 이들 유전자가 각각 ABC (ATP-binding cassette) transporter와hexose transporter를 형성한다는 정도만 밝혀졌다(Krattinger et al. 2009, Moore et al. 2015, Dodds & Lagudah 2016). APR 유전자는 대부분 부분저항성을 나타내기 때문에 다른 APR유전자나 SR 유전자와 조합하게 되면 저항성이 크게 향상된다(Lagudah 2011, Tsilo et al. 2014). 따라서 녹병 저항성 육종프로그램에서는 각 유전자의 저항성 패턴을 정밀하게 평가하고 효과적인 저항성 유전자 2개 이상을 집적(gene pyramiding)하여 저항성 품종을 육성하는 것을 기본 원칙으로 한다(Samsapour et al. 2010, Lowe et al. 2011).
잎녹병 저항성 유전자
잎녹병 저항성 유전자는 Lr로 표기하며 주요 저항성 유전자의 정보는 Table 1에 정리하였다. 약 100년전에 러시아 품종인 Malakof 와 터키 품종인 Webster 에서 Lr1과 Lr2a가 처음으로 보고된 이후 밀의 21개 염색체 모두에서 현재까지 80여개의 잎녹병 저항성 유전자가 보고되었다(Aktar-Uz-Zaman et al. 2017, Qureshi et al. 2018). 잎녹병 저항성 유전자는 대부분 야생밀로부터 일반밀로 도입된 것으로 추정되며 일부는 듀럼밀과 이배체밀에서도 유래한다(Aktar-Uz-Zaman et al. 2017).
잎녹병 저항성 중에 종특이적 저항성은 대부분 유묘기 저항성(SR, seedling resistance)으로, Table 1에서 보는 바와 같이 많은 유전자는 염색체 상의 위치가 밝혀졌고 특정 분자표지와 연관되어 MAS (marker assisted selection)로 육종에 이용이 가능하다(McIntosh et al. 2013, Li et al. 2014). SR 유전자 하나만을 가지고는 저항성을 오래 지속할 수 있는 품종을 만들 수 없는데, 대표적인 SR유전자인 Lr10, Lr14a와 Lr16을 지닌 캐나다 품종 Selkirk의 경우 재배된 지 8년 가량이 지난 후에는 잎녹병에 대한 저항성을 잃게 되었다(McCallum et al. 2016). 또한 SR 유전자 여러 개를 복합적으로 가지는 경우 잎녹병 저항성을 연장시킬 수는 있지만, 다중병원성을 지닌 병원균을 발생시켜 과병원성 집단(hyper-virulent pathogen populations)을 발생시킬 가능성도 높아진다. SR 유전자 중에서는 Lr3, Lr10, Lr20, Lr26를 포함한 소수의 유전자가 육종에 광범위하게 활용되는 편이다(Goyeau et al. 2006).
Slow rusting 계통이 보유하는 APR 유전자는 Lr12, Lr13, Lr22a/b, Lr35, Lr37, Lr48과 Lr49를 포함한 7개는 종특이적이고, Lr12, Lr13과 Lr22b의 경우 종특이적 저항성의 특성인 과민성 세포사 반응(hypersensitive cell death)을 보임에도 불구하고 식물체 발달 후반기에만 저항성을 발현하는 특성이 있다(Singh & Bowden 2011). Lr34, Lr46, Lr67, Lr68, Lr74, Lr75, Lr77과 Lr78은 종비특이적인데, Lr34 (Yr18/Pm38/Sr57), Lr46 (Yr29/ Pm39/Sr58)과 Lr67 (Yr46/Pm46/Sr55)은 흰가루병과 줄녹병, 줄기녹병에도 동시에 저항성을 나타내는 다면발현 특성을 지닌다(William et al. 2003, Spielmeyer et al. 2005, McIntosh et al. 2013, Herrera-Foessel et al. 2014a). 이러한 잎녹병의 APR을 나타내는 품종은 중국 재래종인 Pingyuan 50, CIMMYT에서 선발된 미국 품종 Anza와 멕시코 품종인 Pavon 76이 대표적이다(Li et al. 2014).
다면발현을 나타내는 종비특이적 APR유전자인 Lr34, Lr46과 Lr67에 대해서 좀더 살펴 보면, Lr34의 유전자좌는 7DS로 20세기 초반의 잎녹병 저항성 육종에 많이 활용되었고 환경에 따라 다양한 저항성을 나타내며 앞으로도 50년 이상 저항성을 유지할 수 있을 것으로 기대되고 있다(Krattinger et al. 2009, Lagudah et al. 2009). Lr34유전자는 처음으로 서열이 밝혀진 APR 유전자로, ATP-binding cassette (ABC) transporter단백질을 만들고, 진화적으로 저항성이 감수성보다 늦게 나타났으며 D게놈에 특이적으로 존재하는 것으로 밝혀졌다(Lagudah et al. 2009, Dakouri et al. 2010). Lr46은 멕시코 품종 Pavon 76에서 처음 발견되었으며 CIMMYT 유전자원이 많이 보유하고 있다(Singh et al. 1998, William et al. 2003). Lr46의 분자육종을 위한 마커로는 csLV46이 많이 이용되고 있다(Suenaga et al. 2003, Lillemo et al. 2008). Lr46은 Lr34과 비슷하게 다수의 병원균에 저항성을 보이지만 특정 환경에서는 Lr34 보다는 저항성 정도가 낮다고 보고되었다(William et al. 2003, Lillemo et al. 2008, Kolmer et al. 2018a, Lagudah 2011). Lr67은 파키스탄, 인도와 CIMMYT 유전자원에서 많이 발견된다(Dyck & Samborski 1979, Moore et al. 2015, Hiebert et al. 2010, Lan et al. 2015, Ponce-Molina et al. 2018). Lr67은 hexose transporter 단백질을 만드는데, 저항성(LR67res)과 감수성(LR67sus) 대립인자는 서로 2개의 아미노산만이 다르다(Moore et al. 2015). 저항성인 LR67res 대립인자는 이종이합체화(heterodimerization)를 통해 우성-음성효과(dominant-negative effect)를 나타내는데, 감수성인 LR67sus대립인자에 비해 포도당 친화성이 감소해서 병원균 침입이 억제된다(Moore et al. 2015).
다른 종비특이적 APR 유전자의 특성을 간단히 살펴보면, Lr68 유전자좌는 7BL이고, Lr34과 Lr13를 지닌 브라질 밀 품종인 Frontana에서 최초로 보고되었다(Herrera-Foessel et al. 2012). Lr68 활용을 위한 분자육종에는 7BLNLRR과 csGS가 주로 사용된다(Herrera-Foessel et al. 2012). Lr74 유전자좌는 3BS로, BTSS (BT-Schomburgk Selection)에서 처음 발견되었고 인도 품종 Spark와 미국 품종 Caldwell에서도 보고되었다(Kolmer et al. 2018b). Lr75 유전자좌는 1BS이고, Lr34를 지닌 스위스 품종인 Forno에서 처음 보고되었으며 해당 유전자좌에서 7개 SSR 분자표지를 활용하면 스위스 환경하에서 30% 수준으로 표현형 변이를 설명할 수 있다고 한다(Singla et al. 2017). Lr77 유전자좌는 3BL이고, 미국 품종 Santa Fe에서 보고되으며 SNP 마커인 IWB103가 이용되고 있다(Kolmer et al. 2018c). Lr78 유전자좌는 5DS이고, 브라질 품종인 Toropi에서 동정되었으며 SNP 마커인 IWA6289가 이용되고 있다(Kolmer et al. 2018a). 잎녹병 저항성 강화를 위한 유전자 집적을 위해서는 양적저항성을 지닌 유전자인 Lr16과 Lr23에 수량 감소가 적은 Lr34나 Lr46을 집적하는 방법이 효과적이라고 보고되었다(Stuthman et al. 2007, Tsilo et al. 2014).
녹병은 밀이 순화되던 시기부터 계속해서 생산성과 품질을 위협해온 대표적인 질병으로, 녹병 저항성 밀 품종 육성을 위하여 전세계적으로 많은 연구가 이루어져 왔다. 이를 통하여 현재까지 80여개에 달하는 잎녹병 저항성 유전자가 보고되었고 Lr1, Lr10, Lr21, Lr22a, Lr34/Yr18/Sr57/Pm38, Lr67/Yr46/Sr55/Pm46은 유전자 수준에서 동정이 이루어졌다(Table 1). 많은 수의 잎녹병 저항성 유전자가 보고되었음에도 불구하고 녹병균은 빠르게 진화하여 병원성 패턴을 변화시키므로 새로운 균계가 기존에 저항성으로 알려진 유전자를 붕괴시키는 사례가 지속적으로 발생한다. 따라서 새로운 저항성을 보유한 자원을 탐색하고 저항성 유전자의 염색체상 위치를 결정하기 위한 연구가 지속되고 있으며, 두개 이상의 저항성 유전자를 한 품종에 집적시켜 식물체의 녹병 저항성을 강화하기 위한 연구도 활발하게 진행되고 있다.
국내의 경우 녹병 저항성 품종 개발을 통한 유전적 방제보다는 재배적으로 녹병 발생을 회피하기 위한 전략에 주로 의존해 왔으며, 그간 개발된 밀 품종이나 재래종의 녹병 저항성 여부 및 저항성 유전자 보유에 대한 정보가 미흡하여 관련 연구 문헌을 찾아보기가 어려운 실정이다. 농촌진흥청 국립식량과학원(www.nics.go.kr) 자료에 따르면 1980년대 초까지는 지역적응시험에 공시된 밀 우량 계통의 유묘기 및 성체기 잎녹병 저항성 특성 검정이 실시되었으나, 1980년대 중반 이후에는 이마저도 중단되었다. 국내에 발병하는 잎녹병 저항성 연구를 위해서는 병원균의 수집 및 동정이 매우 중요한데, 잎녹병균인 P. triticina는 활물기생체로 병원균의 유지 관리가 매우 어렵고 농촌진흥청 국립농업유전자원센터(www.genebank.rda.go.kr)에도 등록된 자원이 없는 실정으로, 일부 대학이나 연구소에서 자체 수집한 P. triticina의 여름 포자 균주를 연구에 이용중이다. 이를 이용한 최근 보고에 따르면 국내 육성 밀 40품종에 대한 유묘기 잎녹병 저항성을 기내에서 정밀 평가한 결과 대부분 감수성이었으며, 저항성을 보인 품종은 ‘올밀’과 ‘조농’이 유일하였다(Kim et al. 2019).
국내에서도 기후변화로 인한 이상기온과 국지성 강우 등으로 밀 작기 중에 잎녹병 발생 위험이 점점 증가하고 있다. 이에 대응하여 유전적으로 저항성을 보유한 품종 육성을 통하여 효과적으로 잎녹병 피해를 예방하기 위해서는 앞으로 많은 노력이 필요하다. 우선 국내 환경에서 상품성 또는 적응성이 입증된 기존 품종 및 주요 육성 계통과 재래종에 대하여 잎녹병 저항성을 평가하고 기존에 알려진 저항성 유전자 보유 여부를 판단하여야 한다. 이를 위해서는 잎녹병 저항성 여부를 검증할 수 있는 정밀 스크리닝 체계를 구축해야 하며, 국내 환경을 우점하고 있는 주요 잎녹병 균계를 조사하고 균계 변화를 지속적으로 모니터링하기 위한 연구도 병행되어야 한다. 또한 다양한 잎녹병 저항성 유전자를 국내 밀 품종 개발에 활용할 수 있도록 CIMMYT를 비롯한 국제연구기관 및 주요 국가 연구소와의 국제 공조를 통하여 잎녹병 저항성 유망 유전자원을 적극적으로 도입해야 한다. 분자표지를 활용하여 잎녹병 저항성 대립인자를 국내 품종에 효율적으로 도입하기 위해서는 이미 유전자 수준에서 동정이 이루어져 정밀 분자표지의 활용이 가능한 유전자를 우선적으로 도입하는 것이 효과적일 것이다.
녹병은 전세계적으로 밀 생산성과 품질을 위협하는 대표적 질병으로 유전적으로 저항성을 보유한 품종 육성을 위하여 연구가 활발히 이루어지고 있다. 잎녹병은 Puccinia triticina에 의하여 발생하며 다양한 기후에 적응하므로 발생지역이 가장 광범위하다. 현재까지 80여개의 잎녹병 저항성 유전자가 보고되었고 그 중 6개(Lr1, Lr10, Lr21, Lr22a, Lr34, Lr67)가 동정되었다. 동정된 잎녹병 저항성 유전자 중 Lr34/Yr18/Sr57/Pm38과 Lr67/Yr46/Sr55/Pm46는 다면발현으로 잎녹병, 줄녹병, 줄기녹병과 흰가루병에 동시에 저항성을 보여 특히 주목된다. 우리나라는 아직까지 녹병을 재배적으로 회피하는 수단이 주로 사용되어 저항성 품종 육성을 위한 연구가 미진한 실정이나, 기후변화로 녹병 발생이 증가하고 있어 유전적 저항성 확보를 위한 연구가 시급하다. 우선 국내에서 육성한 품종과 재래종을 대상으로 잎녹병 저항성 여부를 판정하는 한편 해외로부터 저항성 유전자원을 적극 도입하여 저항성 유전자를 다양화해야 한다. 또한 국내 환경에서 주요 균계 분포와 변화를 모니터링함으로써 다양한 저항성 자원을 활용하여 잎녹병 균계 변화에 지속적으로 대응해야 한다. 이와 함께 밀 유전체 분야의 연구를 강화하여 국내 고유의 잎녹병 저항성 유전자 발굴과 분자표지 개발을 병행하여 잎녹병으로 인한 피해를 사전에 예방해야 한다.
본 연구는 농촌진흥청 국립식량과학원 농업과학기술연구개발사업(과제명: 장수형 밀 자원의 농업형질 특성 개선, 과제번호: PJ012792022020)의 지원에 의해 이루어진 것으로 이에 감사드립니다.
Fig. 1
Symptoms of wheat leaf rust disease caused by Puccinia triticina. Photo credit: Dr. Chon-Sik Kang, National Institute of Crop Science, Rural Development Administration.
KJBS-52-213-f1.gif
Table 1
Summary of major leaf rust (Lr) resistance genes.
Table 1
Locus Accz Source Chry Typex Referencesw
Lr1 S79983 Triticum aestivum 5DL SR Qiu et al. 2007, Cloutier et al. 2007
Lr3 - Triticum aestivum 6BL SR Sacco et al. 1998, Herrera-Foessel et al. 2007
Lr9 JK818417 Aegilops umbellulata 6BL SR Schachermayr et al. 1994, Gupta et al. 2005
Lr10 U51330 Triticum aestivum 1AS SR Feuillet et al. 1997, 2003
Lr12 - Triticum aestivum 4BL APR Singh & Bowden 2011, Dieguez et al. 2018
Lr13 - Triticum aestivum 2BS APR Seyfarth et al. 1998, 2000
Lr14a - Triticum aestivum 7BL SR Herrera-Foessel et al. 2008a
Lr15 CB275450 Triticum aestivum 2DS SR Dholakia et al. 2013
Lr16 - Triticum aestivum 2BS SR McCartney et al. 2005
Lr17a - Triticum aestivum 2AS SR Bremenkamp-B. et al. 2008
Lr18 - Triticum timopheevi 5BL SR Sadeghabad et al. 2017
Lr19 FM876846 Agropyron elongatum 7DL SR Gupta et al. 2006, Gennaro et al. 2009
Lr20 - Triticum aestivum 7AL SR Neu et al. 2002
Lr21 AH012974 Triticum tauschii 1DS APR Huang et al. 2003, Neelam et al. 2013
Lr22a KY064064 Triticum tauschii 2DS APR Hiebert et al. 2007, Thind et al. 2017
Lr22b - Triticum aestivum 2DS APR McIntosh et al. 2003, Zhang et al. 2019
Lr23 - Triticum turgidium 2BS SR Chhetri et al. 2017
Lr24 EF011021 Agropyron elongatum 3DL SR Dedryver et al. 1996, Prabhu et al. 2004,
Lr25 - Secale cereale Rosen 4BS APR Procunier et al. 1995, Singh et al. 2012
Lr26 DQ167397 Secale cereale Petkus 1BS SR Mago et al. 2005
Lr27 - Triticum aestivum 3BS,1BL/1RS SR Nelson et al. 1997, Spielmeyer et al. 2003
Lr28 HO772677 Aegilops speltoides 4AL SR Naik et al. 1998, Cherukuri et al. 2005
Lr29 - Agropyron elongatum 7DL APR Procunier et al. 1995
Lr31 - Triticum aestivum 4BL SR Dieguez et al. 2018
Lr32 - Triticum tauschii 3DS APR Autrique et al. 1995
Lr33 - Triticum aestivum 1BL SR Che et al. 2019
Lr34/Yr18/Sr57 FJ436983 Triticum aestivum 7DS APR Lagudah et al. 2009, Krattinger et al. 2009
Lr35 DQ001172 Aegilops speltoides 2B APR Seyfarth et al. 1999
Lr37 AY652952 Aegilops ventricosa 2AS APR Helguera et al. 2003
Lr38 - Agropyron intermedium 6DL APR Mebrate et al. 2008, Yan et al. 2008
Lr39 - Triticum tauschii 2DS APR, SR Raupp et al. 2001
Lr41 - Triticum tauschii 2DS SR Cox et al. 1994, Sun et al. 2009
Lr42 - Triticum tauschii 1DS APR Sun et al. 2010
Lr44 - Triticum aestivum 1BL APR Che et al. 2019
Lr45 - Secale cereale 2AS APR Zhang et al. 2007, Naik et al. 2015
Lr46 - Triticum aestivum 1BL APR Mateos et al. 2006
Lr47 AJ238219 Aegilops speltoides 7AS SR Helguera et al. 2000
Lr48 JG391645 Triticum aestivum 4BL APR Bansal et al. 2008
Lr49 - Triticum aestivum 2AS APR Bansal et al. 2008
Lr50 - Triticum monococcum 2BL SR Brown-Guedira et al. 2003, Blaszczyk et al. 2004
Lr51 AY589012 Aegilops speltoides 1BS APR Helguera et al. 2005
Lr52 - Triticum aestivum 5BS APR Bansal et al. 2011, Qureshi et al. 2017
Lr53 - Triticum dicoccoides 6BS APR Dadkhodaie et al. 2011
Lr56 - Aegilops sharonensis 6AL - Marais et al. 2006, 2010
Lr57 Aegilops geniculata 5DS - Kuraparthy et al. 2009
Lr58 - Triticum aestivum 2BL - Kuraparthy et al. 2011
Lr60 - Triticum aestivum 1DS SR Hiebert et al. 2008
Lr61 - Triticum durum 6BS - Herrera-Foessel et al. 2008b
Lr63 - Triticum monococcum 3AS - Kolmer et al. 2010
Lr64 - Triticum aestivum 6AL SR Kolmer et al. 2019
Lr65 - Triticum aestivum 2AS - Mohler et al. 2012
Lr67/Yr46/Sr55 KR604818 Triticum aestivum 4DL APR Abdelbacki et al. 2014, Moore et al. 2015
Lr68 - Triticum aestivum 7BL APR Herrera-Foessel et al. 2012
Lr70 - Triticum aestivum 5DS - Hiebert et al. 2014
Lr71 - Triticum aestivum 1B - Singh et al. 2013
Lr72 - Triticum durum 7BS SR Herrera-Foessel et al. 2014b
Lr74 - Triticum aestivum 3BS APR McIntosh et al. 2015
Lr75 - Triticum aestivum 1BS APR Singla et al. 2017
Lr77 - Triticum aestivum 3BL APR Kolmer et al. 2018c
Lr78 - Triticum aestivum 5DS APR Kolmer et al. 2018a
Lr79 - Triticum durum 3BL SR Qureshi et al. 2018

zWheat nucleotide accession registered at NCBI (National Center for Biotechnology Information).

yChromosomal location.

xAPR: adult plant resistance, SR: seedling resistance, -: uncertain.

wReference for gene cloning is underlined.

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Research Advances in Wheat Breeding and Genetics for Leaf Rust Resistance
Korean. J. Breed. Sci.. ;52(3):213-224.   Published online September 1, 2020
DOI: https://doi.org/10.9787/KJBS.2020.52.3.213
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Research Advances in Wheat Breeding and Genetics for Leaf Rust Resistance
Korean. J. Breed. Sci.. ;52(3):213-224.   Published online September 1, 2020
DOI: https://doi.org/10.9787/KJBS.2020.52.3.213
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Research Advances in Wheat Breeding and Genetics for Leaf Rust Resistance
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Fig. 1 Symptoms of wheat leaf rust disease caused by Puccinia triticina. Photo credit: Dr. Chon-Sik Kang, National Institute of Crop Science, Rural Development Administration.

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Research Advances in Wheat Breeding and Genetics for Leaf Rust Resistance

Summary of major leaf rust (Lr) resistance genes.

Locus Accz Source Chry Typex Referencesw
Lr1 S79983 Triticum aestivum 5DL SR Qiu et al. 2007, Cloutier et al. 2007
Lr3 - Triticum aestivum 6BL SR Sacco et al. 1998, Herrera-Foessel et al. 2007
Lr9 JK818417 Aegilops umbellulata 6BL SR Schachermayr et al. 1994, Gupta et al. 2005
Lr10 U51330 Triticum aestivum 1AS SR Feuillet et al. 1997, 2003
Lr12 - Triticum aestivum 4BL APR Singh & Bowden 2011, Dieguez et al. 2018
Lr13 - Triticum aestivum 2BS APR Seyfarth et al. 1998, 2000
Lr14a - Triticum aestivum 7BL SR Herrera-Foessel et al. 2008a
Lr15 CB275450 Triticum aestivum 2DS SR Dholakia et al. 2013
Lr16 - Triticum aestivum 2BS SR McCartney et al. 2005
Lr17a - Triticum aestivum 2AS SR Bremenkamp-B. et al. 2008
Lr18 - Triticum timopheevi 5BL SR Sadeghabad et al. 2017
Lr19 FM876846 Agropyron elongatum 7DL SR Gupta et al. 2006, Gennaro et al. 2009
Lr20 - Triticum aestivum 7AL SR Neu et al. 2002
Lr21 AH012974 Triticum tauschii 1DS APR Huang et al. 2003, Neelam et al. 2013
Lr22a KY064064 Triticum tauschii 2DS APR Hiebert et al. 2007, Thind et al. 2017
Lr22b - Triticum aestivum 2DS APR McIntosh et al. 2003, Zhang et al. 2019
Lr23 - Triticum turgidium 2BS SR Chhetri et al. 2017
Lr24 EF011021 Agropyron elongatum 3DL SR Dedryver et al. 1996, Prabhu et al. 2004,
Lr25 - Secale cereale Rosen 4BS APR Procunier et al. 1995, Singh et al. 2012
Lr26 DQ167397 Secale cereale Petkus 1BS SR Mago et al. 2005
Lr27 - Triticum aestivum 3BS,1BL/1RS SR Nelson et al. 1997, Spielmeyer et al. 2003
Lr28 HO772677 Aegilops speltoides 4AL SR Naik et al. 1998, Cherukuri et al. 2005
Lr29 - Agropyron elongatum 7DL APR Procunier et al. 1995
Lr31 - Triticum aestivum 4BL SR Dieguez et al. 2018
Lr32 - Triticum tauschii 3DS APR Autrique et al. 1995
Lr33 - Triticum aestivum 1BL SR Che et al. 2019
Lr34/Yr18/Sr57 FJ436983 Triticum aestivum 7DS APR Lagudah et al. 2009, Krattinger et al. 2009
Lr35 DQ001172 Aegilops speltoides 2B APR Seyfarth et al. 1999
Lr37 AY652952 Aegilops ventricosa 2AS APR Helguera et al. 2003
Lr38 - Agropyron intermedium 6DL APR Mebrate et al. 2008, Yan et al. 2008
Lr39 - Triticum tauschii 2DS APR, SR Raupp et al. 2001
Lr41 - Triticum tauschii 2DS SR Cox et al. 1994, Sun et al. 2009
Lr42 - Triticum tauschii 1DS APR Sun et al. 2010
Lr44 - Triticum aestivum 1BL APR Che et al. 2019
Lr45 - Secale cereale 2AS APR Zhang et al. 2007, Naik et al. 2015
Lr46 - Triticum aestivum 1BL APR Mateos et al. 2006
Lr47 AJ238219 Aegilops speltoides 7AS SR Helguera et al. 2000
Lr48 JG391645 Triticum aestivum 4BL APR Bansal et al. 2008
Lr49 - Triticum aestivum 2AS APR Bansal et al. 2008
Lr50 - Triticum monococcum 2BL SR Brown-Guedira et al. 2003, Blaszczyk et al. 2004
Lr51 AY589012 Aegilops speltoides 1BS APR Helguera et al. 2005
Lr52 - Triticum aestivum 5BS APR Bansal et al. 2011, Qureshi et al. 2017
Lr53 - Triticum dicoccoides 6BS APR Dadkhodaie et al. 2011
Lr56 - Aegilops sharonensis 6AL - Marais et al. 2006, 2010
Lr57 Aegilops geniculata 5DS - Kuraparthy et al. 2009
Lr58 - Triticum aestivum 2BL - Kuraparthy et al. 2011
Lr60 - Triticum aestivum 1DS SR Hiebert et al. 2008
Lr61 - Triticum durum 6BS - Herrera-Foessel et al. 2008b
Lr63 - Triticum monococcum 3AS - Kolmer et al. 2010
Lr64 - Triticum aestivum 6AL SR Kolmer et al. 2019
Lr65 - Triticum aestivum 2AS - Mohler et al. 2012
Lr67/Yr46/Sr55 KR604818 Triticum aestivum 4DL APR Abdelbacki et al. 2014, Moore et al. 2015
Lr68 - Triticum aestivum 7BL APR Herrera-Foessel et al. 2012
Lr70 - Triticum aestivum 5DS - Hiebert et al. 2014
Lr71 - Triticum aestivum 1B - Singh et al. 2013
Lr72 - Triticum durum 7BS SR Herrera-Foessel et al. 2014b
Lr74 - Triticum aestivum 3BS APR McIntosh et al. 2015
Lr75 - Triticum aestivum 1BS APR Singla et al. 2017
Lr77 - Triticum aestivum 3BL APR Kolmer et al. 2018c
Lr78 - Triticum aestivum 5DS APR Kolmer et al. 2018a
Lr79 - Triticum durum 3BL SR Qureshi et al. 2018
Table 1 Summary of major leaf rust (Lr) resistance genes.

Wheat nucleotide accession registered at NCBI (National Center for Biotechnology Information).

Chromosomal location.

APR: adult plant resistance, SR: seedling resistance, -: uncertain.

Reference for gene cloning is underlined.

Table 1

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